Discussion

Unit II dates from 100 to 184 kyr according to ESR dating (see Appendix and Murray et al. 2016) covering Marine Isotope Stages 5 and 6. According to the available dates the studied coprolites are from the very top part of the unit (dated in 100 ± 7 kyr) and belong to Stage 5 (Table 13.1) but the exact age cannot be determined more precisely. They are therefore more likely to represent a warm or a stadial phase than a glacial period.

The present-day river near the area, the terraces of which indicate that it was active in the Pleistocene (Murray et al. 2016), could be a potential source of the diatoms in the one coprolite. Phytoliths in the coprolite 5246 were more corroded than those in 5153 and its associated Unit II deposits, which contained better preserved silica than other deposits. The difference in preservation quality is difficult to explain, except for the indication of guano deposits during and after burial. Corrosion might have resulted from harsh conditions in the surroundings before the phytoliths were accidentally ingested by the animal (as dust), or it might have occurred later under fluctuating water tables or dampness that affected the silica inside the coprolite in the cave. It is known that at present such fluctuations do occur, and it is likely that they also occurred in the past. Corrosion could have been enhanced further by the corrosive qualities of bat guano. Indications of guano and damp/dry fluctuations at the cave interior is indicated by secondary mineral formation, such as tinsleyite, sepiolite, gypsum, ardelite or brushite (Magela da Costa and Rúbia Ribeiro 2001; Marincea et al. 2002; White and Culver 2012) detected by X-ray fluorescence (XRF) and X-ray diffraction (XRD) in both the sediment and the fossils (Marin-Monfort et al. 2016).

Habitat structure inferred through a comparison of the contribution from GSSC phytoliths versus non-grass phytoliths (e.g., Alexandre et al. 1997; Bremond et al. 2005) points to grassy conditions in the region at the time when the coprolites were formed, although the density of woody components cannot be determined. However, the two coprolites differ in content, and the more non-grass inclusions in coprolite 5246 could be related to seasonal factors or could simply be due the possibility that the coprolites represent different habitats in which animals roamed (Fig. 13.4). As is indicated by the non-grass silica like epidermal cells or other round “blocky” phytoliths, several different unidentified plant types could be included in the coprolite assemblages. As can be inferred from the charcoal evidence (Allué 2016) woody species must have occurred locally, especially Prunus. Phytoliths of this genus, which are not produced in fruits, leaves and inflorescences of some species (Kealhofer and Piperno 1998), were not identified, partly because their morphologies are not known (Rovner 1971).

The proportion of GSSC–phytoliths versus indeterminate silica bodies in the coprolite-bearing deposits is similar to that of the Holocene deposits, but the recent soil shows a lower proportion of grasses, which is typical of an overgrazed area like that around the cave at present.

Some are taxonomically and ecologically significant. The underrepresentation of saddle and bilobate phytoliths and comparatively high frequencies of trapezoidal and oblong morphotypes recorded in Unit II clearly suggests that C3 grassy conditions prevailed at the time when the coprolites were formed and the place where they were ingested outside the cave. This is supported by the presence of polylobates recorded throughout Unit II. Polylobates are recorded in at least 25% (n = 31) of modern Pooid species (Rossouw 2009). The phytoliths also indicate the presence of other plants which can at present not be identified.

The surrounding area at the time of the coprolite production could also have undergone dry summers resembling that of alpine meadows of the Crimean Mountains or that of the cold-dry steppe (winter-rain) of South Jordan (Cordova 2011). Because of the presence of Stipa-type in an area where Paniceae and Danthonioideae are rare, the occurrence of grasses of the Stipeae tribe, most of which reflect cold and dry continental climates, is suggested.

The coprolite phytolith assemblages only give a reflection of what is available in the environment and not necessarily of the actual proportions of plant types. Potential bias in ratio towards more grasses in the GSSC in relation to unidentified silica in the coprolite samples is plausible in view of possible selective consumption of grasses by carnivores as is recorded in ecological studies worldwide (Skinner 1976; de Arruda Bueno et al. 2002). However, comparison with the phytoliths in the surrounding deposits of Unit II does not suggest any marked bias. The cave deposit samples from the Unit II sediment may be a more unbiased reflection of the vegetation in the immediate surroundings than the coprolite because they do not favor behavioral selection from a wider range.

The oblong/trapezoid phytolith ratios between the coprolites and surrounding deposits differ slightly with more oblong types in the latter. This could be from widely roaming animals trapping phytoliths in their dung and not from the local slopes next to the cave (as represented by the cave deposits). In comparison to present conditions as reflected by the modern sample outside the cave, oblong types are more prominent but it is not possible to say if this is due to climate a different climate or modern grazing disturbance. The assemblages that occur in Unit I deposits during the Holocene compare well with those in Unit II, suggesting that climates did not differ markedly.

On the basis of other evidence the habitat varied (Andrews et al. 2016). The large mammals and charcoal indicate deciduous woodland while small mammals,

amphibians and reptiles indicate open steppe environments. The taphonomy of the latter group suggests that they were probably brought to the cave from a distance by predators in a setting similar to the present, where woodland occurs in the vicinity of the cave and steppe not too far away. Therefore it is not impossible that woodland existed similar to the vegetation that can potentially develop in the area today under current climatic conditions and no agricultural disturbance.

Conclusions

1.Pollen was extremely rare in the two carnivore coprolites investigated, and none was found in the sediments. The lack of pollen is probably due to environmental conditions and the location of the excavation 40 m into the Azokh 1 passageway.

2.Phytoliths were abundant in the coprolites and in the deposits of Unit II. Nine different grass silica short cell (GSSC) phytolith types were identified, and these indicate that the vegetation type was most likely a temperate C3-grass steppe mosaic.

3.Phytoliths other than those of grasses were recorded and they could have been derived from local woodland. Caution is needed with the interpretation of the openness of the vegetation in view of the unknown degree of possible selection of phytoliths by the carnivore and due to the limitation that a large number of the phytoliths were not identified.

4.The few diatoms recovered suggest the availability of local water.

5.Long silica structures (longer than 200 microns) were observed in one of the coprolites. They resemble sponge spicules and indicate wet conditions.

6.The discovery of numerous phytoliths show that the Azokh deposits have great potential for a phytolith study and interpreting environmental conditions throughout the whole Azokh sequence. A more detailed analysis can therefore be undertaken beyond the scope of this study. The potential is demonstrated in deposits at the older Dmanisi site in the Georgian Caucasus that contain comparable phytolith assemblages indicating marked changes in water stress in the region (Messager et al. 2010).

Acknowledgments We thank Yolanda Fernández-Jalvo for providing the coprolites, initiating the study and providing relevant information. We are also grateful to the authorities of Nagorno-Karabakh for the support and permissions to work on these specimens. We are grateful to Tania King and diggers for careful work collecting these fossils, as well

as field assistants for modern soil sampling on the slope of the cave. Thanks are extended to Karen Hardy for collecting sediment samples from the section of Azokh.

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